Proceedings of the XI International Symposium on Biological Control of Weeds
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New research on Alternanthera philoxeroides (alligator weed) in its South American native range Alejandro J. Sosa,1 Mic H. Julien2 and Hugo A. Cordo1 Summary Alternanthera philoxeroides (alligator weed) is a herbaceous amphibious weed of the Amaranthaceae, native to southern South America. Several agents from Argentina, e.g. Agasicles hygrophila and Arcola malloi, have been used to control aquatic A. philoxeroides in Australia and the USA. However, in Australia, the weed continues to pose a serious problem, particularly in terrestrial situations. In Argentina, A. philoxeroides is distributed along the catchments of the Paraná and Uruguay rivers in the north, and in the catchments of the San Borombón and Salado rivers in the centre of Buenos Aires prov- ince. Two forms are recognized: A. philoxeroides f. philoxeroides in the southern range and A. philoxeroides f. angustifolia in the northern range. There appears to be preferential attack by flea beetles on A. philoxeroides f. angustifolia. In 2000, the CSIRO initiated a collaborative research project with the USDA South American Biological Control Laboratory in Argentina to search for new biolog- ical agents. After the initial year of surveys, the natural enemies that may have biological control poten- tial included: two species of leaf-feeding beetles, Systena spp.; a tip-galling Cecidomyiidae fly; and two agromyzid flies, one that causes node galls and another that mines leaves. Two fungi were also found: one probably Nimbya alternantherae, known to have a wide host range, and another, thought to be a new Sphaceloma species, that causes a characteristic “corky” deformation on the stem and leaf surfaces. Surveys will be extended and the interactions between these herbivores and pathogens with A. philoxeroides will be studied. Keywords: alligator weed, Alternanthera philoxeroides, native range, natural enemies. Introduction moth were subsequently released in Australia and controlled the weed in warm temperate aquatic habitats In its introduced range, Alternanthera philoxeroides (Julien 1981). However, terrestrial growth of A. (Martius) Grisebach (alligator weed; Amaranthaceae) philoxeroides and aquatic growth in cooler regions of is often a serious aquatic and terrestrial weed. Surveys Australia continue to cause serious concern (Julien & for natural enemies were carried out in the 1960s in Bourne 1988, Julien & Stanley 1999). parts of its native range in South America (Vogt 1961), Alternanthera philoxeroides consists of several taxa and the flea beetle Agasicles hygrophila Selman and in both its native and adventive ranges. It was first Vogt, the moth Arcola malloi (Pastrana) and the thrips described by Martius in 1826, and named Bucholzia Amyinothrips andersoni O’Neil were released in the philoxeroides. Covas (1939, 1941) considered two vari- USA. Good control was obtained in aquatic habitats, eties of alligator weed: A. philoxeroides var. obstusifolia largely attributed to the flea beetle and the moth (Moquin) Hicken and A. philoxeroides var. acutifolia (Spencer & Coulson 1976). The flea beetle and the (Moq.) Hicken. The former was characterized by the presence of ovate lanceolate leaves with obtuse or sub obtuse apex; whereas the latter variety, acutifolia, has 1 South American Biological Control Laboratory, USDA-ARS, Bolivar lanceolate leaves with acute apex. Pedersen (1967) indi- 1559 (B1686EFA), Hurlingham, Buenos Aires, Argentina. cated that Martius did not consider two varieties of alli- 2 CSIRO Entomology, Long Pocket Laboratories, 120 Meiers Road, Indooroopilly, Queensland 4068, Australia. gator weed and he considered that the specimen Corresponding author: A.J. Sosa <[email protected]>. deposited in Brussels, which had ovate or elliptic and 180 New research on alligator weed in South America obtuse leaves, as a good lectotype. However, Pedersen located from Posadas, on the northern Argentina border (1999) suggested that A. philoxeroides be divided into with Paraguay, to Tandil, well south of Buenos Aires. two and he referred to these as forms: A. philoxeroides f. In the field, the presence of fruits and seedlings of A. philoxeroides (Mart.) Griseb. and A. philoxeroides f. philoxeroides was recorded, and collected for cultiva- angustifolia Süssenguth. tion in the laboratory. Seedlings and stem cuttings were Records from the United States suggest that both collected from four localities and also grown in the forms were introduced there. Ganstad & Solymosy laboratory. Two localities represented the southern (1973) and Weldon et al. (1973) reported that some form, Tandil and Mar del Plata in Buenos Aires prov- plants of A. philoxeroides had small stems, and in some ince, and two represented the northern form, Santa Fé, cases the internodes were solid, and apparently this Santa Fe province, and Hurlingham, Buenos Aires tended to produce a deficient plant that is not preferred province. After 6 months of growth in identical condi- by the flea beetle A. hygrophila. Kay & Haller (1982) tions, 27 plant parameters were measured and found two different forms in the USA, which they compared using principal components analysis. Three differentiated as the narrow-stemmed alligator weed factors were extracted that explained 71% of the total (NSA) biotype and the broader-stemmed alligator weed variance. They were: diameter of internode of leaf one, (BSA) biotype. The NSA biotype was characterized by length/width ratio of leaf one, and leaf apical angle of the presence of slender stems with short internodes and leaf one. ANOVA was carried out to evaluate mean obtuse and rounded leaves. The BSA biotype has differences and Tukey test was used for multiple broader and longer stems and longer and acute leaves. comparisons among pairs of means based on unequal They pointed out a different pattern in the damage sample of sizes. All statistical analysis was carried out caused by A. hygrophila. Populations of NSA seem to using Statistica 5.5. be attacked less than the BSA populations in the USA and both biotypes responded differently to herbicides (Kay 1992). Wain et al. (1984) demonstrated genetic Results differences between the two biotypes in the USA using isozyme pattern analyses. In contrast, no genetic varia- Alligator weed: native ranges, tion was detected within or between populations of A. morphological variation and biology philoxeroides in China using RAPD analysis (Xu et al. Alternanthera philoxeroides f. philoxeroides, the 2003). southern form, was distributed along the catchments of In 2001, the CSIRO (Australia) initiated a coopera- San Borombón and Salado rivers in Buenos Aires prov- tive research project with the USDA South American ince (Fig. 1). It was not found in western Buenos Aires Biological Control Laboratory in Argentina to update province. It was also found in some sites in the north the list of natural enemies known from A. philoxeroides west of Argentina, in township drains, possible outside and to identify potential new biological control agents. the native range. This form has small, ovate leaves, short internodes and slender stems. It occurred in semi- Materials and methods aquatic conditions, in ditches next to roads or along the shores of lakes. Flowering was abundant in summer, Surveys were conducted, mostly in Argentina, but also with short, small inflorescences. in Uruguay, Paraguay and south-eastern Brazil, The second, northern form, A. philoxeroides f. between October 2001 and November 2002. Natural angustifolia, was distributed along the Paraná River enemies were sampled at 93 A. philoxeroides sites. from Posadas and along the Paraguay River from Sites where Alternanthera aquatica (Parodi) Chodat Pantanal region (Brazil). It also occurred along the (ex A. hassleriana) occurred were also sampled. At Uruguay River downstream from Santo Tomé (Corri- each A. philoxeroides site, adult insects were collected entes province). Thus, it extended from the northern either by direct aspiration from plants or after sweeping parts of Buenos Aires province through the north-east with a net, placed in 70% ethanol and sent to taxono- wetlands of Argentina (including Entre Rios and Corri- mists for identification. Other immature insects were entes province, western Misiones and eastern parts of collected alive and reared in the laboratory to adult the provinces that have their eastern borders along the stage. These were also sent for identification. Paraná and Paraguay rivers), and along the south and The extent of the native ranges of both forms of A. east coast of Uruguay and possibly into south-east philoxeroides were approximated using the local distribu- coastal areas of Brazil. This form was also found in the tion of the weed, its morphology, and assessments of north-west of Argentina and at one site in the north of where it grew, i.e. natural areas or highly disturbed loca- Patagonia in Río Negro province (Fig. 1). In these cases tions such as town drains. Herbarium specimens from they were only associated with human activities and representative sites, six stems per site, were collected. For were considered to be outside the native range. Vogt et each of the six stems, two younger leaves were removed al. (1979) suggested that north-western populations and stored in 96% alcohol for genetic analysis. RAPD were isolated relicts or introduced populations. We analyses were conducted on material from seven sites suggest the latter. 181 Proceedings of the XI International Symposium on Biological Control of Weeds 70° 65° 60° 55° 50° 45° 40° 15° 15° Native